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(Issued 4th July, 1962) 


The Application of the Generic Name Macropus Shaw 1790 and of Other 
Names Commonly Referred to the Grey Kangaroo J. H. Calaby, 

George Mack and W. D. L. Ride .25 

A New Species of Thylacoleo and Notes on Some Caudal Vertebrae of 

Palorchestes azael . . Alan Bartholomai . 33 

Fossil Marsupials and Cainozoic Continental Stratigraphy in Australia: A 

Review J. T. Woods .41 


J. H. Calaby, Wildlife Survey Section, C.S.I.R.O., Canberra 
George Mack, Queensland Museum, Brisbane 
W. D. L. Ride, Western Australian Museum, Perth 

There has been considerable discussion in recent years as to the identity 
of the kangaroos collected in the vicinity of the Endeavour River, north-east 
Queensland in 1770 by Captain James Cook’s party. Three specimens were obtained, 
weighing respectively 8, 38 and 84 pounds. From an account of this material, 
Muller (1776) described the kangaroo ( Mus canguru) mentioning only the 38 pound 
specimen. This is the holotype of Mus canguru Muller and it has been generally 
accepted that the specimen was a grey kangaroo. 

Iredale and Troughton (1925), as a result of examining the description 
contained in Solander’s unpublished manuscript (1768-1771), expressed doubt that 
the original specimen described and named Mus canguru by Muller (1776) was a grey 
kangaroo. They suggested that it might have been a wallaroo of the robustus group. 
Later, the same authors (1937) endeavoured to show that it was really a whiptail 
or pretty- faced wallaby ( Wallabia elegans). Again, this conclusion was based on 
Solander’s manuscript, aided by the purchase of two skins collected near Cooktown 
on the Endeavour River, one a whiptail wallaby and the other an antilopine wallaroo. 
It is difficult to understand this decision. Solander’s description is a composite one ; 
parts of it (one of the weights, some measurements, a sex) can clearly be related to 
the holotype, while other parts have been taken from other specimens. 

Raven (1939) then discussed the matter and held that the earlier revisers 
were correct in indentifying the first described specimen as a grey kangaroo. Morrison- 
Scott and Sawyer (1950) provided excellent support for this view, and they produced 
additional pertinent evidence that the type of Mus canguru Muller was a young 
grey kangaroo. These authors published (1) two outline sketches (indeterminable*) 
of an entire kangaroo made by Parkinson, artist on board the Endeavour, (2) wash 

* We have examined a coloured transparency (supplied through the Mitchell Library, 
Sydney) of a painting of a kangaroo by Stubbs which is probably the original of the figure in 
Hawkesworth (see Lysaght 1957). We are unable to identify it specifically with any species 
which nowadays occurs at Cooktown. Any attempt to establish that this figure and painting 
are of the holotype would be pointless. Muller specifically nominated the holotype by reference 
to its weight and there is no evidence that this figure represents the nominated animal. 




drawings by Nathaniel Dance of a skull, which can be identified as that of Macropus 
robustus, a grey wallaroo ; and (3) a photograph of a skull of a young grey kangaroo 
which they designated the “ photo- lectotype ” of Macropus canguru (Muller). This 
last skull was collected on Cook’s voyage and was given to John Hunter by Banks, 
a member of Cook’s party. It is No. 1732 in Owen’s catalogue (1853) and No. 3703 
in Flower’s catalogue (1884). It is the skull of a young grey kangaroo and we believe 
that it is from the 38 pound specimen collected near the Endeavour River. Both 
Owen and Flower described this skull and its dentition. 

Morrison -Scott and Sawyer have suggested that the largest of the three 
specimens collected, weighing 84 pounds, was a wallaroo of the Macropus robustus 
group, and now represented by the Dance drawings. The presence of a specimen 
of the wallaroo in Cook’s collection would account for the statement by Solander in 
his composite description that the internarial area of the kangaroo was naked. 
A juvenile grey kangaroo from Cook’s voyage (presumably the specimen which weighed 
8 pounds) was recorded by Gray (1843) as being preserved in spirits in the British 
Museum. It is no longer in the collections. The skull of the third animal, which was 
in the collections of the Royal College of Surgeons in London, was destroyed by 
a bomb and is now represented by a photograph in the British Museum and 
undoubtedly it is the skull of a young grey kangaroo, shown by us here to be from 
an animal weighing in the vicinity of 38 pounds, and certainly not 84 pounds. 

The instability which now exists in the nomenclature of these well known 
kangaroos and wallabies is undesirable and can best be removed by proving the 
specific identity of the holotype which was nominated by Muller. Clearly, no 
consideration need be given to the 8 pound juvenile grey kangaroo which was in the 
British Museum ; the holotype is either the Hunterian specimen or another, probably 
represented by the specimen drawn by Dance. It must be emphasised that Solander’s 
composite description which was not published until 1925 is of no nomenclatural 


Morrison-Scott and Sawyer lacked material from the type locality, and with 
this in mind, a party from the Queensland Museum, consisting of D. P. Vernon, 
S. Breeden and M. E. McAnna, collected in the vicinity of Cooktown, Endeavour 
River, during October and November, 1960. Among the specimens obtained are 
five grey kangaroos, seven grey wallaroos, six antilopine wallaroos, seven whiptail 
wallabies and ten agile wallabies. 

A young male grey kangaroo (J.10749), collected at Kings Plains, November 
24, 1960, by Vernon and Breeden, is nearest in weight to the 38 pound animal of 
the Endeavour party. Although it weighed 55 pounds, including viscera and stomach 
contents, its dental age is precisely that of the 38 pound animal. Another male 



(J. 10750), weighing 90 pounds, including viscera and stomach contents, was collected 
at the same locality. This specimen has all molars fully erupted, and not only has 
dP 4 been shed, but its successor (P 4 ) also has been shed on the right side and is close 
to being shed on the left. It will be clear from these remarks that the 84 pound 
Endeavour specimen cannot have been the skull given by Banks to Hunter, and on 
the evidence afforded by the specimen (J. 10749) in the Queensland Museum, it must 
have been the 38 pound animal. Furthermore, in addition to agreeing exactly in 
dental age, the limb measurements of the young male (J. 10749) alone agree closely 
with those provided by Muller when describing Mus canguru. 

The exact stage of dental eruption reached by the Hunterian (38 pound) 
specimen was well described by both Owen and Flower. P 3 has been shed, dP 4 was 
about to be replaced by P 4 ; M 1 , M 2 and M 3 were in place in the maxilla and M 4 was 
still in its crypt. The dentition of J. 10749 is the same as will be seen from the 
accompanying plates. 

It was not possible to establish on the recent Cooktown material the range 
of variation in weight which can be expected at this stage of dentition. However, 
for this purpose, Mr. W. H. Butler, Associate of the Western Australian Museum, 
collected a series of male grey kangaroos from a single population at Congelin, 
south-west Australia. Three of these were at the same dental age as both of the 
above, and their weights were 63, 68 and 79 pounds, a weight range of 16 pounds. 
The south-west grey kangaroo is a heavier animal than the north-east Queensland 
form, but the difference in minimum and maximum weights is similar to the difference 
between the 38 pound specimen of Cook’s party and the Queensland Museum 
example, J. 10749. It would appear that the 17 pounds difference is not significant. 

Finally, in order to satisfy ourselves as to the probability of the identification 
of the skull of the Dance drawing with the 84 pound specimen, this (as reproduced 
in Morrison -Scott and Sawyer) has been carefully compared with skulls from the 
Endeavour River of the whiptail wallaby, the agile wallaby, the antilopine wallaroo, 
the grey wallaroo and the grey kangaroo, and it is clearly a specimen of the grey 
wallaroo. At first sight, the third upper incisor appears unusual until it is realised that 
Dance, in order to show the presence of the faint groo\e in the outer surface of the 
posterior lobe, has slightly overemphasized it. The skull drawn by Dance has a 
sectorial tooth followed by three fully erupted molariform teeth, and a partly open 
alveolus is shown behind the last molariform tooth. The cementum area of the root 
of the third incisor is well exposed and the pronounced supraorbital crests become 
confluent above the temporal fossa to form a sagittal crest. These growth characters 
support our identification of the teeth as P 4 , M 1 , M 2 , and M 3 (with M 4 in its crypt). 
Hawkesworth says of the 84 pound animal, that it was “ not at its full growth, the 
innermost grinders not yet being formed.” A male specimen of the grey wallaroo 
at this dental age (J. 10738) was collected at Annan River, 17 miles south of Cooktown 
and it weighed 70 pounds. 



The skull of the wallaroo drawn by Dance is very probably that of the 
84 pound animal shot by Lieutenant Gore on July 27th, 1770. Its presence in the 
series would explain the presence of the characters which are atypical of the grey 
kangaroo in the composite description of Solander (e.g. the naked internarial region). 
The grey wallaroo is the only <c grey ” macropod in the area which possesses these 
characters and at the same time achieves a weight of 84 pounds. Further it should 
be noted that, of the five species collected at the Endeavour River in 1960, the 
male grey wallaroo is the only male form which is sufficiently nondescript externally 
to be included with a series of grey kangaroos without remark. Cook’s party obtained 
two grey kangaroos, and male specimens of species other than the grey wallaroo are 
either widely different from the grey in colour or are, even to the untrained eye, 
brightly and obviously ornamented. 


It has already been stated that the final decision of Iredale and Troughton 
that Mus canguru Muller was based on a whiptail wallaby was not derived from an 
examination of Muller’s description, but from an examination of a description 
contained in a manuscript by Solander, now in the British Museum. This description 
included both the male and female genitalia and the pouch, and the weights of 
three animals were given. The main characters stated by Solander and used by 
Iredale and Troughton were that the area between the nares was naked and that 
the third incisor was bilobed, broad from side to side and with smaller anterior 
lobes. As Morrison-Scott and Sawyer have pointed out, Solander’s description 
of the incisors accords neither with the grey kangaroo nor the whiptail wallaby, 
but it is in accord with the condition in the wallaroo. 

Solander also stated “ Par intimum Molarium diu intra alveolus sous latit, 
in junioribus non discernandum,” implying that both the 38 and 84 pound specimens 
had unerupted last molars. He referred to the 38 pound example as a male of two 
or three years, and to the 84 pound animal as “ adultus.” Similarly, Hawkesworth 
described the 38 pound animal as “ a young one, much under its full growth. ” 
Elsewhere in the same work Hawkesworth made it clear that the term “ full growth ” 
referred to the possession of fully erupted molar teeth since he said of the 84 pound 
animal “ We found that this animal was not at its full growth, the innermost 
grinders not being yet formed.” 

All male whiptail wallabies between 30 and 50 pounds in weight collected 
on the recent field trip to the Endeavour River had erupted last molars, as was to be 
expected. Externally, each individual has a prominent white stripe, approximately 



12 mm. wide, extending from the nares to behind the eye on each side of the face ; 
another prominent white stripe, crescent-shaped, on each thigh ; and the fur under 
the head, centre of throat and entire abdomen is white. It is most unlikely that those 
who examined and described the first collected Macropodidae from Australia would 
ignore these vivid markings if the whiptail wallaby was represented in the collection. 


From the above evidence, together with the evidence provided by Morrison- 
Scott and Sawyer, it would appear that the following specimens were collected in the 
vicinity of the Endeavour River, north-east Queensland, in July, 1770 by Cook’s 

(1) A grey wallaroo, Macropus robustus, weighing 84 pounds. It is likely 

that from this specimen Solander in manuscript described the area 
between the nares as naked. Now apparently represented by a wash 
drawing of the skull by Nathaniel Dance in the British Museum. 

(2) A young grey kangaroo, weighing 38 pounds. This specimen, which 

had P 4 and M 4 still in their crypts, was described by Hawkesworth 
(1773) and described and named Mus canguru by Muller in 1776. 
Now represented by a photograph of the skull in the British Museum. 

(3) A juvenile specimen, weighing 8 pounds, a grey kangaroo. Recorded 

as preserved in the British Museum (Gray, 1843), but not now in the 


Macropus is currently the generic name of the large-sized kangaroos, including 
the grey kangaroo, and it has been applied in this way for more than one hundred 
years. The genotype is Yerboa gigantea Zimmermann (1777) which is accepted as 
a synonym of Mus canguru Muller (1776). 

The view of Iredale and Troughton is that the type of Mus canguru Muller 
was a whiptail wallaby, at present generally known as Wallabia elegans. To those 
who accept this view, the generic name Macropus must be used for the large-sized 
wallabies, and another name will have to be found for the large-sized kangaroos. 



Since the case presented here rests, to some extent, on probability, in order 
to stabilize both the generic and specific names involved, we have decided to approach 
the International Commission for Zoological Nomenclature with the following 
request : — 

(1) That the specimen, the skull of which was given by Sir Joseph Banks 
to John Hunter and became No. 3703 in the collections of the Royal 
College of Surgeons, London, be declared the holotype of Mus canguru 
Muller 1776 ; that since this specimen has been destroyed, it be 
replaced by a neotype, a grey kangaroo, Queensland Museum 
No. J.10749, male, skin and skull, collected at Kings Plains, 20 miles 
south of the Endeavour River, November 24, 1960, by D. P. Vernon 
and S. Breeden. 

(2) That a procedure be adopted to make the name Yerboa gigantea 
Zimmermann 1777 an objective synonym of Mus canguru in accordance 
with the current usage of these two names. 

(3) That the name Macropus major Shaw 1800 be conserved for the grey 
kangaroo with type locality Sydney as restricted by Iredale and 
Troughton (1934). According to Article 7 2d of the International Code, 
Macropus major has the same type specimen as Yerboa gigantea and 
the Plenary Powers will have to be exercised in order to validate the 
action of Iredale and Troughton. 

If the Commission accepts this recommendation, the generic name Macropus 
will continue to be available for the large-sized kangaroos ; M . canguru will be the 
name of the grey kangaroo ; and major will be available for the grey kangaroo of 
Sydney and beyond should it prove to be subspecifically distinct from the Endeavour 
River, north-east Queensland grey kangaroo. 

It is a pleasure to acknowledge the assistance rendered by the staff of the 
Mammal Department, British Museum (Natural History), in particular Miss Jean 
Ingles and Dr. Gordon Corbet ; the Fisheries and Wildlife Department, Victoria, 
especially Mr. John McNally ; Mr. E. H. M. Ealey of the Department of Zoology 
and Comparative Physiology, Monash University ; and Mr. W. H. Butler, Associate 
of the Western Australian Museum. 




"Flower, W. H., 1884. Catalogue of specimens ... in the Museum of the Royal College 
of Surgeons of England, 2, p. 708. 

Gray, J. E., 1843. List of . . Mammalia in the British Museum, p. 87. 

Hawkesworth, J., 1773. An account of the Voyages undertaken . . . for making Discoveries 

in the Southern Hemisphere ... by Captain Cook, 3, p. 577 (1st Ed.), p. 173 
(2nd Ed.). London. 

Iredale, T., and Troughton, E.LeG., 1925. Captain Cook’s Kangaroo. Av&t. Zool., 3, p. 311. 

, 1934. A Checklist of the Mammals Recorded from Australia, p. 54. 

, 1937. The identity of Cook’s Kangaroo. Rec. Aust. Mus,, 20, p. 67. 

Lysaght, A., 1957. A clue to the mystery of Captain Cook’s Kangaroo. The New Scientist, 3, p. 17. 

Morrison- Scott, T. C. S., and Sawyer, F. C., 1950. The identity of Captain Cook’s kangaroo. 
Bull. Brit. Mus. (Nat. Hist.), Zocl., 1, p. 43. 

Muller, P. L. S., 1776. Syst. Nat. (Linne), Suppl., p. 62. 

Owen, R., 1853. Descriptive Catalogue of the Osteological Series . . . Museum 

Royal College of Surgeons of England, 1, p. 322. London. 

Raven, H. C., 1939. The identity of Captain Cook’s Kangaroo, J. Mammal ., 20, p. 50. 

Shaw, G., 1800. Gon. Zool., 1, pt. 2, p. 505. 

-Zimmermann, E. A. W., 1777. Specimen zoologiae geographicae, Quadrupedum domicilia et 
migrationes sisteus & etc., p. 526. Leyden. 

Macropus canguru. Grey Kangaroo 
Plate V. Doral view of skull. 

Plate VI. Ventral view of skull. 
Plate VII. Lateral view of skull. 
Plate VIII. Mandible. 

All views of same skull, J. 10749, male (Q.M.). All natural size 
Specimen from Kings Plains, 20 miles south of Cooktown, N.E. Queensland. 


mm m 

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Oueemland Museum J~l Q74-L> 


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Ouecmlrtod Museum 



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Alan Bartholomai 
Queensland Museum 

The Darling Downs area of south-eastern Queensland consists mainly of 
extensive fluviatile deposits of Upper Cainozoic Age. Woods (1960) indicated that 
the Chinchilla Sand, developed in the valley of the Condamine River in the north- 
western Darling Downs, was probably deposited earlier than the Pleistocene alluvia 
to the south-east. 

Specimens of the genus Thylacoleo Owen in the collections of the Queensland 
Museum were revised by Woods (1956) and those from the south-eastern Darling 
Downs were attributed to T. carnifex Owen. A small proportion of fragmentary 
material from the Chinchilla district was tentatively separated as possessing features 
sufficiently distinct to constitute a possible new species. Recently, a partial right 
mandibular ramus was collected from the Chinchilla Sand at the Chinchilla Riflo 
Range (Rifle Range Number 78, parish of Chinchilla), supporting the view that the 
variations are characteristic, and the material is here described as a new species. 

All measurements are in millimetres. 


(Figures 1, 2) 

Material. F.3565, holotype. A partial right mandibular ramus with l t 
broken, P 3 - M 2 , young adult. Chinchilla Sand, at 363677 Chinchilla 4 mile military 
map, possibly Pliocene. 

Specimens from the Chinchilla Sand at Chinchilla, north-western Darling 
Downs : F.2957, partial right mandibular ramus, no teeth preserved. F.296R 

partial left mandibular ramus with both P 3 and M x broken, very aged. F.2962, 
partial right mandibular ramus with P 3 , adult. F.2960, partial left mandibular ramus 
with M 2 , aged. F.2964, partial left mandibular ramus with l ± broken, P 3 , juvenile. 
F.2963, broken left P 3 , adult. F.2495, partial left mandibular ramus with P 3 , adult. 
F. 2941, partial right maxilla with both P 3 and M 1 broken, adult. F.2954, partial 
right maxilla with both P 3 and M 1 broken, adult. F.2955, partial right maxilla with P 3 ,. 



Specimens from the Darling Downs : F.3569, partial right mandibular ramus 
with both 1 1 and P 3 broken, M 1? adult. F.3570, partial left mandibular ramus with 
Mj, adult. F.3571, partial left mandibular ramus. F.3572, partial left mandibular 
ramus with both P ;j and M x broken, aged. 

Measurements of Mandibles 


Length of 
crown of 

Breadth of P 3 

posterior root 

Length of 
crown of Mi 

Breadth below 
of M t 

Angle between 
1 1 and base 
of mandible 

Holotype, F.3565 
















































Ramus deep, strong, particularly robust in region of implantation of cheek teeth ; 
longitudinal axis very slightly convex laterally ; symphysis unfused ; symphysial plane short, 
deep, subequilaterally triangular, somewhat upturned. Fossa subalveolaris deep, confluent ; 
mental foramen prominent, ventral to anterior margin of P 3 and antero-dorsal to junction of 
-anterior margin and inferior surface of ramus at blunt angle ; lateral alveolar walls of P 3 thinning, 
roots becoming exposed with age, with simultaneous development of inter-rootial depression. 
Ramus ascending at low angle posterior to smooth, weak diagastric process ventral to M 2 ; 
postalveolar ridge not prominent, ascending gradually posteriorly, disappearing on mesial wall 
of large coronoid process. Process diverging from line of ramus, directed antero-posteriorly ; 
antero-dorsal margin ascending at approximately 45°, grooved, laterally flanged ; flange continuing 
ventrally on body of ramus limiting large, anteriorly deep, ectocoronoid fossa. Wall of fossa 
perforated by masseteric foramen opening into inferior dental canal close to mandibular foramen. 
Condyle, postero-mesial angle of ramus and posterior portion of coronoid process not preserved. 

Lower median incisor broken, but enough remains to indicate the presence of shallow, 
posterior longitudinal furrows ; lingual furrow best developed. 1 x making an angle of approxi- 
mately 42° with base of mandible. P x and P 2 not retained ; alveoli small, mesiad to anterior 
root P 3 . Third premolar elongate, sectorial, very convex laterally, deeply rooted ; roots directed 
somewhat dorso-laterally ; base of crown swollen ; enamel finely ridged vertically, particularly 
•on lingual surface, thickened to flange on anterior edge ; crown broadest above posterior root, 
not tapered posteriorly, asymmetric, with labial face set at much higher angle than lingual ; 
prominent antero-lingual buttress associated with anterior cuspid. Main surface of wear near 
planar, developed ventro- lateral to cutting edge. Enamel also removed labially at base of crown. 
Molar series reduced. Mj relatively large, subtriangular ; anteriorly with metaconid high, flanged, 
with short longitudinal cutting edge in functional continuity with that of P 3 ; base of crown broad 
-anteriorly due to development of prominent labial buttress ; crown posteriorly reduced with low 
median ridged area and shallow, finely ridged, dorso- labial fossette. Facet of wear developed 



in continuation with that of P 3 , and another directed postero-laterally ; birooted, anterior root: 
much larger than posterior. M a small, with single root partially divided lingually, not functional;, 
crown with shallow, faintly ridged dorsal depression. 

Figure 1. Thylacoleo crassidentatus sp. nov. F.3565, holotype. Right ramus, lateral and occlusal? 
views. Natural size. 

Woods (1956) compared the fragmentary Thylacoleo material from Chinchilla 
with T. carnifex from the south-eastern Darling Downs and discussed the differences 
exhibited. Although only slight dissimilarities are observed in the maxillary remains,, 
the mandibles differ considerably. 

The most significant feature is the structure of the cheek teeth and, in-, 
particular, that of the lower sectorial premolar, where the tooth is posteriorly much 
broader and the longitudinal convexity much stronger than in the corresponding P 3 . 
of T. carnifex. M x exhibits a difference in relative proportions ; it is comparatively 
stout owing to the greater width above both the anterior and posterior roots. The 
posterior portion of the crown is comparable with the size of the posterior root and. 



-exhibits a much wider median ridged area together with the development of a shallow, 
postero- labial fossette. The second molar, although variable developed, is stronger 
than in T. carnifex and its alveolus is partially divided by a vertical ridge on the 
lateral wall. In one specimen (F.2960), a depression immediately posterior to the 
relatively large second molar is interpreted as the alveolus of a very small third molar. 

In addition, the ramus is much wider in the region of implantation of the cheek 
deeth and displays a less pronounced longitudinal concavity of the mesial side, while 
.a more recumbent lower median incisor is indicated by the reduction of the angle 
between the incisor and the base of the mandible. 

Figure 2. Thylacoleo crassidentatus sp. nov. Lateral and occlusal views of maxillary fragment 
(F.2954). Natural size. 



Owing to the fragmentary nature of the preservation of the maxillary material 
in T. crassidentatus relatively few comparative measurements are possible. The third 
premolars exhibit no features which differ from those of T. carnifex. The base of the 
crown of the first molar in F.2954 is wider than in T. carnifex, while in F.2955 the 
tooth is more distinctly tri-rooted, the anterior root being largest, while the posterior 
roots are subequal. In addition, F.2954 exhibits a depression immediately posterior 
to M 1 and this is considered to represent the alveolus of a small second molar. 

It is apparent that the molar series of T. crassidentatus are much less reduced 
both in size and numbers than in T. carnifex, a fact in keeping with the suggested time 
relationships of the two faunas. 

(Figures 3, 4) 

Six caudal vertebrae (F.3564), were recently located in position in the alluvia 
of “ Strathmore ” station near Collinsville, north-eastern Queensland, and were 
presented to the Queensland Museum by Mr. E. Cunningham. Associated with the 
series is a large number of post-cranial fragments together with an incomplete right 
mandibular ramus, the lower median incisors, and several maxillary fragments of 
JPalorchestes azael. In view of the field association there can be little doubt that the 
vertebrae, interpreted as representing the first six of the caudal series, belong to 
this species. 

Neural arches and zygopophyses are not preserved in first, fourth and fifth 
vertebrae, while transverse processes are broken in vertebrae five and six, and are 
poorly represented in the first. No chevrons are preserved. 


Vertebrae large, stout. All centra short antero-posteriorly, broader posteriorly than long 
and broadest in second. Length decreases from first to the fifth, then increases. Centra broadly 
convex interiorly ; posterior depth greatest in first caudal vertebra, decreasing regularly to sixth. 
Epiphyses firmly fused. Floors of neural canals in anterior vertebrae subdivided by median 
ridges, on either side of which foramina pierce centra ; distal centra pierced by one foramen, with 
median ridges discontinuous. Neural arches moderately high and broad with neural spine quite 
high in second caudal, decreasing in development posteriorly and barely visible as low ridge in 
■sixth vertebra. Zygopophyses functional in proximal vertebrae but with anterior zygopophyses 
reduced and functionless by sixth vertebra and with posterior zygopophyses lost. Transverse 



processes broadly expanded antero-posteriorly in anterior vertebrae, particularly at extremities, 
but are rounded at tips ; processes decrease in transverse diameter and in antero-posterior 
expansion to sixth caudal. 

Figure 3. Palorchestes azael Owen. Dorsal view of caudal vertebrae (F.3564). One third natural 




Caudal Vertebra 

Length of 

breadth of 

depth of 

Breadth across 































In his revision of the species of Palorchestes Owen from south-eastern Queens- 
land, Woods (1958) concluded that the genus belongs within the family Diprotodontidae 
and not within the Macropodidae as indicated by Owen and later workers. The 
erroneous idea of the systematic position of Palorchestes led Owen to attribute a number 
of post-cranial macropodid remains to the genus. Fletcher (1945) mentioned additional 
post-cranial fossils referred to P. azael in the collections of the Australian Museum, 
Sydney, but Woods doubted the identification because of the absence of field 
association. Woods has also indicated that the lower incisor found associated with 
post-cranial remains from the dune sandstone at Fowler’s Cove, Nepean Peninsula, 
Victoria and tentatively referred by Gregory (1902) to this species, is of characteristic 
macropodid aspect and referable to one of the large extinct species of Protemnodon 

Relatively few of the Australian fossil diprotodontids have their caudal 
vertebral series sufficiently well known to afford comparisons with those of P. azael. 
The posterior breadths of the proximal centra in P. azael and Diprotodon optatus Owen 
exceed the lengths, but while the centra of D. optatus are shallow posteriorly owing 
to the flattening of the inferior surfaces, those of P. azael are similar in structure to 
the macropodids. Gill and Banks (1956) and Scott (1915), show that the centra of 
Nototherium tasmanicum Scott are structurally similar to those of D. optatus. The 
transverse expansion of the transverse processes and the rounding of the process 
extremities in P. azael is similar to that of the other diprotodontids, and is especially 
similar to that of N. tasmanicum. The anterior neural canals, however, are comparable 
with those observed in the Macropodidae, being high and comparatively broad, but 
contrasting markedly with the low, very broad neural canals in D. optatus. In 
conjunction with this feature, the zygopophyses in the proximal caudal vertebrae 
of D. optatus are much reduced and functionless, while the neural spines are 
represented by extremely low tubercles. This is similar to the structure observed 
in N. tasmanicum but is in direct contrast to the well-developed, functional 
zygopophyses and moderately high neural spines in P. azael. 



Figure 4. Palorchestes azael Owen. Anterior view of second caudal vertebra (F.3564). One third 
natural size. 

Tedford (1959) recorded a palorchestine diprotodontid from the Etadunna 
Formation of possible Oligocene Age (Stirton et al., 1961), at Lake Ngapakaldi and 
Lake Kanunka North, Tirari Desert, north-eastern South Australia, and noted that 
the tail was long and heavy. 

The palorchestine diprotodontids were apparently lightly built, more mobile 
grazing animals than other members of the Diprotodontidae. Although the tail of 
P. azael differs considerably from that of D. optatus, it does appear to be structurally 
closer to that of N. tasmanicum. The structural similarities to the macropodid tail 
are believed to be related solely to body form and are not of general systematic 


Fletcher, H. O., 1945. Palorchestes — Australia’s Extinct Giant Kangaroo. Aust. Mus. Mag., 8, 
pp. 361-365. 

Gill, E. D., and Banks, M. R., 1956. Cainozoic History of Mowbray Swamp and Other Areas 
of North-western Tasmania. Rec. Q. Viet. Mus., 6 (n.s.), pp. 1-42. 

Gregory, J. W., 1902. Some remains of an extinct kangaroo in the Dune-Rock of the Sorrento- 
Peninsula, Victoria. Proc. Roy. Soc. Viet., 14 (n.s.), pp. 139-144. 

Scott, H. H., 1915. A Monograph of Nototherium tasmanicum. Rec. Oeol. Surv. Tasm., 4,pp. 1-47. 

Stirton, R. A., Tedford, R. H., and Miller, A. H., 1961. Cenozoic Stratigraphy and Vertebrate- 
Paleontology of the Tirari Desert, South Australia. .Rec. S. Aust. Mus., 14, pp. 19-61. 

Tedford, R. H., 1959. Notes. Soc. Ver . Paleont. News Bull., No. 55, pp. 23-26. 

Woods, J. T., 1956. The Skull of Thylacoleo carnifex. Mem. Qld. Mus., 13, pp. 125-140. 

, 1958. The Extinct Marsupial Genus Palorchestes Owen. Mem. Qld. Mus., 13, 

pp. 177-193. 

— , 1960. Fossiliferous Fluviatile and Cave Deposits. Jour. Geol. Soc. Aust., 7, 

pp. 393-403. 



J. T. Woods 

Geological Survey of Queensland, Brisbane 

The geological history of the mammals in other parts of the world suggests 
that marsupials entered the Australian Region and became isolated at an early date, 
at least by the beginning of the Cainozoic era. However, prior to the discoveries of 
the last decade, the early palaeontological record of Australian marsupials was most 
unimpressive, with only one species, Wynyardia bassiana, known from lower Tertiary 
deposits. Their abundance in Quaternary sediments stood in remarkable contrast. 

Since then, the work of Professor R. A. Stirton and his colleagues in the 
eastern part of the Lake Eyre Basin in South Australia has provided a major 
contribution to the palaeontological record of the marsupials and other vertebrates 
in Tertiary time. Other significant discoveries have been reported from the Tertiary 
of Victoria, where the association of some of the fossils with marine beds has enabled 
more precise dating of the remains. While the bulk of our knowledge is still confined 
to upper Cainozoic forms, their potential value in continental stratigraphy is 
apparent. At the same time, the variety of these upper Cainozoic marsupials points 
to the diversification of the groups in this continent at an early date, and emphasizes 
that the paucity of the early Tertiary record will have to be overcome before the 
basic phylogeny of these groups can be established. 


The oldest Tertiary marsupial, the age of which can be stated with any 
precision, is still Wynyardia bassiana from marine sediments at Fossil Blulf near 
Wynyard, northern Tasmania. The skeleton was originally found in a fallen block 
of limestone from the “ Turritella Bed,” and the results of the application of the 
fluorine test, published by Gill (1957), appear to establish that this bed was its true 
provenance. On the basis of its correlation with the Janjukian Stage in Victoria, 
an Oligocene age is now generally accepted for the marine section at Fossil Blulf. 
Following his re-examination of the skeleton, Wood Jones (1931) concluded that the 
affinities of Wynyardia were with the Phalangeridae. 

Several fragmentary marsupial fossils have been recovered from shallow 
water marine beds and from horizons interbedded with marine sediments in Victoria. 
As our knowledge of such forms increases, their occurrence will provide a basis for 
correlation of the continental Tertiaries with the marine succession. The oldest of 



these remains is portion of a macropodid femur reported by Glaessner, McGowran,. 
and Wade (1960) from sand of Balcombian age (Middle Miocene) above the Bochara 
Limestone in Grange Burn, near Hamilton, Victoria. Younger marine sediments in 
the same area, the Grange Burn Coquina of lower Pliocene age, have yielded a 
fragment of a macropodid mandible, referred by Stirton (1957b) to the subfamily 

Three diprotodontid fossils originally found on the beach at Beaumaris, 
Victoria, and described by Stirton (1957b) were shown by Gill (1957) to have a 
fluorine index comparable with a provenance in the Black Rock Member of the 
Sandringham Sand. These marsupials then belong to the Cheltenham “ Stage ” of 
upper Miocene age. 

Details of the stratigraphic succession and additional information on the fauna 
of the Cainozoic sediments of the Tirari Desert in the eastern part of the Lake Eyre 
Basin, South Australia, have been recently published by Stirton, Tedford, and 
Miller (1961). While the stratigraphic relationships of the various units have been 
established through superposition, the authors have expressed difficulty in assigning; 
them ages in terms of the conventional epochs of the Tertiary. The oldest unit 
recognised, the lacustrine Etadunna Formation, is tentatively assigned to the 
Oligocene on the basis of comparative evolutionary studies of the macropodid fauna. 
The formation postdates the development of duricrust on representatives of the 
Cretaceous Winton Formation and remnants of early Cainozoic fluviatile deposits. 

It must be acknowledged that lateritic processes were operative not only over 
wide areas in Australia, but at more than one time during the Cainozoic. However,. 
Twidale (1956) has postulated that in north-west Queensland uplift and dissection 
of a widespread lateritized surface, developed on Cretaceous and probable early 
Tertiary rocks, took place approximately in Miocene time. The surface was apparently 
of considerable extent in inland Australia, and constitutes the Australian Pediplain 
of King (1950), the erosion of which, he claims, was initiated by earth movements in 
late Oligocene or Miocene time. It may well be that a tentative Miocene age for 
the Etadunna Formation would be more appropriate than a tentative Oligocene age. 

The marsupials of the Etadunna Formation recorded by Stirton et al. (1961)< 
include a dasyurid, a phascolarctid, Perikoala, previously described by Stirton 
(1957a), two macropodids, and a small diprotodontid with possible palorchestine- 

Lithologically the formation is calcareous in part, with some massive limestones 
carrying chert nodules. Paten (1961) has recently discussed the Cainozoic freshwater 
limestones and associated sediments, which are widespread in western Queensland,, 
but which show their greatest development in the valleys of the Georgina and Burke 
Rivers and near Birdsville, all within the present Lake Eyre drainage basin. Their 


section usually includes detritus from older lateritized profiles at the base, and silicifica- 
tion frequently occurs towards the surface. Paten has suggested a late Tertiary or 
early Quaternary age for these deposits. Vertebrate fossils have been recovered from 
only one locality, in the Carl Creek Limestone, near Riversleigh in the Gregory River 
valley. This is away from the main occurrences, and the fragmental bones, so far 
recovered, are unsatisfactory for close study. 

It is possible that deposition of calcareous lacustrine sediments took place 
at different times in the Cainozoic over this whole belt. No correlation of any of these 
deposits with the Etadunna Formation can be suggested on the available evidence ; 
there is no detailed lithological resemblance, but the similarity in gross stratigraphic 
relationships indicates that the possibility might be considered in future work. 

In the Tirari Desert the next unit recognised by Stirton et al. (1961), the 
Mampuwordu Sands, consist of fluviatile sediments deposited disconformably or 
possibly unconformably on representatives of the Etadunna Formation. Elements 
of the fauna of the stream channel deposits, tentatively placed as lower Pliocene in 
age, were described by Stirton (1955). In the recent work the marsupials of the 
faunal list now comprise the peramelid Ischnodon, four macropodids including 
Prionotemnus , and two diprotodontids, Meniscolophus and a form with affinities to 

Unfossiliferous sandy and argillaceous sediments of the Tirari Formation, 
which overlie the Etadunna Formation unconformably, are also tentatively referred 
by these authors to the Pliocene. Fossiliferous Pleistocene sediments, the Katipiri 
Sands, and younger Quaternary or Recent fluviatile and aeolian sediments complete 
the succession in the Tirari Desert, the most complete recognised in continental 
deposits of the Cainozoic of Australia. 

To date, elements of the fauna of the Mampuwordu Sands have not been 
found elsewhere in Australia. Stirton (1955) suggested that Nototherium watutense, 
originally described by Anderson (1937) from the Watut River, New Guinea, was 
probably referable to Meniscolophus. Information from Dow (1961, personal 
communication) indicates that the New Guinea species occurs in sediments of 
Pleistocene age, but some occurrences may be older. 

Loosely compacted fluviatile and lacustrine sediments which are tentatively 
referred to the Pliocene are widespread in Queensland. They comprise the Glendower 
Formation and its lithological equivalents, which are mostly unnamed and unmapped, 
although the Lynd Formation is known to be extensive in the plains east of the 
southern part of the Gulf of Carpentaria. Conglomerates containing pebbles of the 
silicified duricrust, commonly known as “ billy,” are usually prominent in the 
section, and the sediments appear to have been derived from the dissection of the 
extensively lateritized, peneplaned (or pediplaned) middle Cainozoic surface. They, 



in turn, often display broad ferruginous mottling, usually interpreted as a weak 
lateritic effect. There is evidence of broad warping, and in places considerable 
dissection has followed uplift. 

Unfortunately these widespread deposits are usually unfossiliferous, but the 
prevalence of leached non-calcareous elastics lessens the chances of preservation of 
vertebrate remains. An exception is the Chinchilla Sand (Woods, 1960) which has 
been traced on the basis of both lithology and fossil content for a distance of nearly 
40 miles in its surface and subsurface extent between Warra and Nan gram Lagoon 
in the north-west of the Darling Downs. The maximum thickness is about 100 feet 
and calcareous horizons occur. Tortoise and crocodile remains are abundant as well 
as marsupials in the extensive vertebrate fauna. There is a striking difference 
between the diprotondontid elements of this fauna and those of the superficial 
deposits of the eastern Downs. Of these Euryzygoma is absent from the presumably 
younger alluvia, while Euowenia and Palorchestes are represented by distinct species. 
However superposition has yet to be established. 

A few vertebrate fossils are known from deep alluvia, beneath basalts, in 
southern Australia. Some of these occurrences are probably upper Tertiary, such 
as those at Buninyong, Victoria, at a depth of 238 feet (Gill, 1957), and the Canadian 
Lead at Gulgong, New South Wales, at a depth of 130 feet (Dun, 1895). Freshwater 
sediments are known to occur below the upper Cainozoic volcanics of north Queensland, 
but only plant remains have been recovered from them. 

Additional localities listed by Gill (1957) for the possible occurrence of Tertiary 
marsupials in southern Australia include One Tree Point, Hobart, and the Geilston 
Travertine in Tasmania, the site at Smeaton, Victoria, whence came the dasyurid, 
Glaucodon ballaratensis Stirton, and the lacustrine deposits at Coimadai, Victoria. 


Vertebrate remains assigned to the Quaternary are widespread in fiuviatile, 
lacustrine, and cave deposits, while there are more restricted occurrences in spring 
deposits, aeolianites, and tuffs. Detailed stratigraphic studies have yet to be made 
on many of these, and no precise correlation of the various faunas can be made. 
Indeed, the most comprehensive attempt at a post-Tertiary chronology for Australia 
is still that of Browne (1945). 

While the fragmentary evidence of the upper Tertiary marsupial remains 
indicates that the major groups were differentiated in this continent by the Miocene, 
many of them reached their acme in the Pleistocene. The fluctuating climate of 
that epoch and the consequential rapid changes in the environment no doubt 
maintained strong selection pressure, especially on browsing and grazing herbivores 


of the open forests and grasslands. The possible results, rapid evolution, with 
increasing specialization, gigantism, and extinction, are all evident in the- 
palaeontological record of the Quaternary. 

The occurrence of any fossil marsupials to the north of the Australian mainland, 
is of particular interest, in that information may be gained on the times and directions 
of dispersal of the various marsupial groups. To date, their occurrence is restricted 
to the diprotodontid, Nototherium watutense , and a few fragmentary undescribed 
macropodids from the Morobe Goldfields area. Information kindly supplied by 
Dow (1961, pers. commun.) indicates that these fossils are derived from the Otibanda 
Lake Beds which were deposited in two lakes in the valleys of the Bulolo and Watut 
Rivers, separated by extensive andesitic volcanics of the Bulolo Gorge. The sediments 
of the smaller area near Wau comprise conglomerate, sandstone, siltstone, and mud- 
stone, with interbedded andesitic tuff and agglomerate near the base. A Pleistocene 
age is considered likely for this sequence. The larger lake, in the valleys of the 
Watut and lower Bulolo, was formed by faulting and regional uplift in the Snake 
River area. The presence of andesitic volcanic material, apparently subaerially 
deposited, throughout much of the section, suggests that in part this sequence may 
be slightly older. The Otibanda Lake Beds are deformed, with dips up to 45°. 

In a small collection recently received from this area the only well preserved 
specimen is portion of a diprotodontid mandible referable to Nototherium sp. Its 
molar pattern is very similar to Nototherium tasmanicum from the upper Pleistocene^ 
of Tasmania and the more widely distributed N. mitchelli of the Pleistocene. However, 
this comparison does not assist correlation since the isolated upper premolar figured 
by Stirton (1957b) suggests that the genus Nototherium ranged at least from upper 
Miocene time. 

One of the most intensively collected areas of Pleistocene alluvia in Australia 
occurs in the valleys of the Condamine River and its tributaries of the eastern 
Darling Downs. Many of the type specimens of the Pleistocene marsupials described 
by Sir Richard Owen in the last century were obtained from these deposits. In the 
eastern part of the area, as in the valley of King Creek, the fossiliferous beds comprise 
brown calcareous clays with lenticular basaltic gravels at rather shallow depths in the 
creek sections. Along the Condamine River near Dalby and Macalister, the observed 
sections are thicker and commonly contain sands and grits as well as brown and grey 
clays. Bore records show up to 167 feet of alluvia in this area. A general Pleistocene 
age has been assigned to this sequence by Woods (1960) based on the almost complete 
absence of living species among the fossils. It is feasible that the more superficial 
fossiliferous alluvia along King Creek are upper Pleistocene in age, while the thicker 
sediments in the Condamine valley range back to include equivalents of the Chinchilla 



Marsupials are predominant in this fauna from the eastern Darling Downs ; 
reptiles are not so common as in the Chinchilla Sand. The most common of the giant 
marsupials is the widespread Diprotodon optatus, and the dominant marcopodids 
are Macropus titan and Protemnodon anak. The smaller polyprotodont marsupials 
.and the phalangerids are not well represented, but those with a general forest habitat 
and small size are unlikely to be preserved in the fluviatile deposits of wide valleys 
and plains. 

Following the early work of Owen, a large number of fossil marsupials were 
described from the Darling Downs by C. W. De Vis. His descriptions are frequently 
unaccompanied by locality and stratigraphic data, and in some cases type specimens 
were not designated. This applies, in particular, to his work on the macropodids 
(De Vis, 1895), which deals in composite fashion with collections from both the 
Chinchilla area and the eastern Darling Downs, and stratigraphic evaluation of many 
of his species has not been possible. Currently, revisionary work on these forms is 
being undertaken by Dr. W. D. L. Hide and Mr. A. Bartholomai. 

Diprotodon optatus was widely distributed on the mainland, and while it 
Teached King Island, there is no record of its having reached Tasmania. It was 
apparently adapted to a wide range of habitats ; Gill (1955) records its occurrence 
at altitudes between sea level and 2,000 feet. The apparent absence of Diprotodon 
from Tasmania may be due to a relatively late dispersal of the genus and the 
presence of an indigenous species of Notothcrium ( N . tasmanicum) as well as the wide- 
spread N. mitchelli in the Mowbray Swamp Peat, regarded by Gill and Banks (1956) 
as upper Pleistocene, may be taken as evidence in support of this view. While the 
restriction of the range of the family Diprotodontidae in southern Australia to the 
upper Pleistocene or the upper part of the middle Pleistocene as suggested by Keble 
(1945) is no longer tenable, indications are that most occurrences of Diprotodon 
optatus in this region are in upper Pleistocene deposits. Furthermore, the species 
is known to range into early Recent time. 

However, Diprotodon sp. occurs in the Chinchilla Sand, and Owen (1870) 
recorded the genus from a depth of 100 feet in the Condamine alluvia. Its apparent 
variation in time range may reflect the local patterns of sedimentation in different 
parts of the continent. 

Two separate faunas have been recognised by Stirton et al. (1961) from the 
Katipiri Sands of Pleistocene age in the Tirari Desert. These fluviatile deposits 
rest disconformably on the Tirari Formation. The presumably older mammalian 
fauna from Lake Kanunka, tentatively referred by these authors to the early 
Pleistocene, contains not only marsupials but the oldest rodent known from the 
Australian Region. The marsupials comprise a dasyurid, a thylacoleonid, two 
phascolomids including one specimen referable to Phascolonus, several macropodids 
■of diverse groups, and diprototodontid fragments, possibly referable to Euowenia. 
Diprotodon has not been collected in this fauna. 


The other mammalian fauna, placed as late Pleistocene, includes several 
rodents, as well as marsupials, comprising the dasyurid Sarcophilus, the phalangerid 
Trichosurus, the phascolomid Phascolonus, several macropodids including the still- 
living Bettongia lesueuri, and the widespread diprotodontid Diprotodon. 

A large assemblage of living and extinct marsupial genera, including Diprotodon, 
has been obtained from the lowest unit of the aeolian sequence at the archaeological 
site at Lake Menindee (Tedford, 1955). This fauna was contemporaneous with 
aboriginal man with the Tartangan culture, and radiocarbon dating of the site 
shows it to be of early Recent age at approximately 6,570 years before the present 
time (Tindale, 1957). People with the Tartangan culture and those with the earlier 
Kartan culture both reached Tasmania, so no zoogeographic basis for the failure of 
Diprotodon to reach there is apparent. 

Few of the extinct marsupials seemed to have survived the time of the Mid- 
Recent Thermal Maximum, about 5,000 years ago. The macropodid Procoptodon 
survived until the time of the Pirrian culture of australoid people, about 4,250 years 
ago (Tindale, 1957). It would appear that the extinction of Pleistocene marsupial 
species was progressive, not a catastrophic result of any sudden climatic change,, 
although the demands of the fluctuating enviroment on genetically senile populations 
were certainly important factors. The arrival of Man probably had a critical effect 
on certain species, especially those of small numbers in restricted areas. 

Cave earths and tufas have yielded abundant fossil mammals, including 
many not known or poorly represented in other deposits ; but these differences 
largely reflect differences in habitat and mode of accumulation of the remains. 
Predators, including owls, appear responsible for most of the material. Native rodents 
and bats are well represented in some occurrences. The assemblages usually comprise 
a mixture of extinct and living species. While many of the deposits are of upper 
Pleistocene or even younger age, it is possible that some collections represent material 
from more than one stratigraphic horizon. Furthermore, the time ranges of many 
living species of small marsupials have not been established, and the taxonomic 
evaluation of the fragmentary material often presents difficulties. 

Notes on the cave fossils of south-western Western Australia were published 
by Glauert (1948) and these embody references to his earlier work in the area. 
A radiocarbon date of > 37,000 years for one of these occurrences, the Mammoth 
Cave, has been indicated by Ride (1960). In this latter paper the fauna of the 
Wombeyan Caves, New South Wales, is discussed and regarded as upper Pleistocene 
in age. 

Studies of cave faunas of Recent age in southern Western Australia by 
Lundelius (1957) show that the geographic ranges of many living species were vastly 
different in prehistoric time. Very recent material from caves in the Buchan district 



■of Victoria, listed by Wakefield (1960), is of particular interest in that it includes 
Burramys parvus, previously known only from an assemblage of living and extinct 
species from the Wombeyan Caves. 

As biostratigraphic studies involving our Quaternary mammals proceed, it will 
be necessary to give special consideration to the local time ranges of species, which 
vary with time of dispersal and time of extinction. This variation becomes of greater 
significance with the decrease in age of the fauna. 

Mr. Duncan Dow, of the Bureau of Mineral Resources, kindly furnished 
information on the stratigraphy of the Otibanda Lake Beds, in the Morobe Goldfields 
-area, Territory of New Guinea. 


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S. G. Reid, Government Printer, Brisbane